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Primary Production in the Columbia River Estuary I. Spatial and Temporal Variability of Properties!

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Primary Production in the Columbia River Estuary I. Spatial and Temporal Variability of Properties! Pacific Science (1990), vol. 44, no. 1: 17-37 © 1990 by University of Hawaii Press. All rights reserved Primary Production in the Columbia River Estuary I. Spatial and Temporal Variability of Properties! J. RUBEN LARA-LARA,z BRUCE E. FREY,3,4 AND LAWRENCE F. SMALL3 ABSTRACT: Light , major nutrients, water temperature, turbidity and its or­ ganic and inorganic fractions, chlorophyll, phaeophytin, DCMU [3-(3,4­ dichlorophenyl)-l, I dimethyl ureal-enhanced fluorescence (DCMU ratio), par­ ticulate organic carbon (POC), particulate organic nitrogen (PON), and primary production were measured from April 1980 through April 1981 in a 65-km stretch ofthe Columbia River estuary. Daily solar input, light attenuation in the water, and chlorophyll concentration accounted for 75% of the variability of daily primary production in the main estuarine axis and 85% in the shallows. The rapid appearance of a turbidity load created by the Mt. Saint Helens volcanic eruption in May 1980and the subsequent clearing of the water as the load moved out ofthe estuary became a natural experiment to show that light availability was indeed the limiting factor to phytoplankton production in the estuary. Spatial variability in chlorophyll concentration was caused mainly by large summer reductions at the location where freshwater cells were lysed on contact with low­ salinity intrusions. Mean values for properties in the main axis generally were not significantly different from those in the shallows, suggesting that the main axis and shallows experience similar , rapid flushing times. Total primary production for the estuary was almost 30,000 metric tons C yr", but areal production was only 100 g C m- 2 yr" , which puts the Columbia system at the low end ofNorth American estuaries. The low areal production was likely a result of light limita­ tion, chlorophyll reduction at the low-salinity boundary, and a short residence time of water and viable cells in the estuary. OF ALL THE aquatic ecosystems, estuarine sys­ adapt. Factors vary in response to tidal, diel, tems are perhaps the ones exhibiting the widest and seasonal cycles, as well as to sporadic and most frequent physicochemical fluctua­ changes caused by storms and mankind's in­ tions and the ones to which phytoplankton tervention. For these reasons, understanding (and other organisms) find it most difficult to of the mechanisms controlling the estuarine abundance of phytoplankton, its production, and the species composition ofpopulations is most difficult. Yet estuaries are of primary I This study was supported by a grant from the Pacific Northwest River Basins Commission and the U.S. Water importance both from an ecological and an Resources Council, administered through the Columbia economic view.The lower Columbia River is a River Estuary Data Development Program (CREDDP) coastal plain estuary, or drowned river valley, and Columbia River Estuary Study Taskforce (CREST) . separating the states of Washington and J. R. Lara-Lara had a scholarship from the National Oregon in northwestern USA. The estuary Council of Science and Technology in Mexico. Manu­ script accepted 12 May 1989. was formed as the sea level rose to its present 2Centro de Investigaci6n Cientifica y Educacion Su­ position after the last glaciation. However, the perior de Ensenada , Ave. Espinoza no. 843, Ensenada, Columbia River estuary is much more river­ B. c., Mexico 22830. dominated than most other coastal plain estu­ 3College of Oceanography, Oregon State University, Corvallis, Oregon 97331. aries in the world. The Columbia River is the 4 Present address: Oregon Health Sciences University, second longest river in North America, with Portland, Oregon 97201. the second largest volume of discharge in 17 18 PACIFIC SCIENCE, Volume 44, January 1990 FIGURE I. Map of the study area. Estuarine regions (numbers) are delineated by hydrographic and sedimentary properties. Zones (letters) are (a) marine zone. (b) estuarine mixing zone, and (c) freshwater, or tidal fluvial. zone. the United States (annual discharge is about mainly on river hydrology, circulatory pro­ 2 x 1011 m - 3 , or 58% that of the Mississippi cesses, and sedimentary geology, Simenstad et River). The Columbia and its tributaries drain al. (in press) sectioned the estuary into three 2 a region of ca. 660,500 m • River flow is now principal zones and 10 different regions for greatly regulated by dams, but still varies from purposes of analysis along the estuarine con­ about 3,000 to 12,000 m' sec-I. tinuum and in the adjacent bays. For our Williams and Scott (1962) and L. G . Williams phytoplankton studies, confined to the water (1964, 1972) reported the phytoplankton flora column, we could not distinguish between re­ of the estuary. Haertel and Osterberg (1967) gions 3 and 5 in the estuarine mixing zone, and and Haertel et al. (1969) described some as­ therefore treated that area as combined region pects of hydrography and plankton ecology (3 + 5). for this system, and Park et al. (1969, 1972) Nine cruises were conducted approximately discussed the estuarine nutrient budget. How­ every other month from April 1980 to July ever, before our research there had been no 1981. Stations in both shallows and main reports on phytoplankton production for this channels were sampled (Figure 2). With ex­ ecosystem, and no attempts to understand the ception of the June and July 1981 cruises, mechanisms that control the phytoplankton when only three stations were sampled, the biomass in the estuary. Here we report the number of primary production experiments spatial and temporal distribution of phyto­ per crui se varied from 5 to 12. To study the plankton biomass and primary production as spatial and temporal variability of tempera­ well as the distributions of the physicochemi­ ture, selected inorganic nutrients, particulate cal driving variables in the estuary throughout chlorophyll a and phaeophytin a, particulate an annual cycle. organic carbon and nitrogen, total suspended particle load, and the organic and inorganic fractions ofthe total suspended load, from 25 to 47 stations were sampled on each cruise. MATERIALS AND METHODS All stations were sampled at the surface, and The study area extended from the mouth of at the deeper stations depths of 2.5, 5.0, and the estuary 56 km inland (Figure 1). Based 10.0 m were sampled with Van Dorn bottles. Primary Production in the Columbia River Estuary I -LARA-LARA ET AL. 19 FIGURE 2. Sampling stations for the investigation of water column primary production. 0 = biomass measurements only; • = primary production and biomass measurements. The cross-hatched areas are those containing the shallow­ water stations . Primary production was assessed by the puted from alkalinity measurements made by radiocarbon uptake method (Strickland and potentiometric titration with 0.1 N H 2S04 , as Parsons 1972). Because the shallow euphotic described by Wetzel and Likens (1979). zone was always well mixed, samples were all Because phytoplankton was distributed collected from the surface. Once collected, the homogeneously through the photic zone, and samples were labeled with carbon-14 and in­ because the light intensity production rate cubated in 80-ml clear polycarbonate bottles, relationship was unchanging over the ex­ with two replicates for each incubation. Incu­ perimental time (Lara-Lara 1982), conversion bations usually were done for 4 hr around of primary production during the incuba­ local noon under natural sunlight in clear deck tion period to daily integral production 1 tanks. Surface water was circulated through (mgC m-2 d- ) was reasonably straightforward the tanks to maintain temperature. Except for (Vollenweider 1971). Basically, a "surface rate surface samples, light was attenuated with curve" was determined from an equation neutral screens to 50, 30, 15,6, 1, and 0% of describing the photosynthesis: depth curve incident light. At the end of all incubations, (Vollenweider 1971) and a curve ofdaily pho­ the radiolabeled samples were immediately fil­ tosynthetically active radiation (PAR). The tered through 0.8-/lm-pore-size membrane fil­ ratio of the area under this surface rate curve ters, and the filters were preserved in Aquasol to that fraction of the curve during which in individual liquid scintillation vials. No sig­ incubations were made was used to convert to 1 nificant differences were found when we com­ mg C m- 2 d- . Specification of the photosyn­ pared fumed (HCl) and unfumed carbon-14 thesis: depth curve and daily PAR curve re­ samples. This was expected, because the estu­ quired measurement of PAR at the estuary arine phytoplankton was mainly diatoms and surface (10 ) and determination of the diffuse microflagellates (Amspoker and McIntire light extinction coefficient (k). 1986). Radioactivity was analyzed in a Beck­ Daily PAR (295-695 nm) was measured by man Model 7500 liquid scintillation counter. an Epply precision recording pyranometer Carbonate alkalinity, for conversion of radio­ mounted at a land-based location about activity to carbon-based production, was com- 30 km up-estuary from the mouth. Light pen- 20 PACIFIC SCIENCE, Volume 44, January 1990 etration into the water column, for determina­ data (Jay and Smith [in press]; Sherwood and tion ofk and the photic depth, was measured Creager [in press]) were selected to represent with a Licor submersible spherical quantum distributions ofproperties along the main axis meter. of the estuary, from near the mouth through Chlorophyll a and phaeophytin a analyses, the most riverine
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